Forensic Science International: Reports 1 (2019) 100046

Contents lists available at ScienceDirect

Forensic Science International: Reports

journal homepage: www.elsevier.com/locate/fsir

Novel application of the ‘Hardy-Weinberg equilibrium’ in the analysis of

cusp patterning phenotype in Gujarati Population
Jayasankar P. Pillai a, * , Dhwani Patel b
a
Institute of Forensic Sciences, Gujarat Forensic Sciences University, Gandhinagar, India
b
Division of Forensic Odontology, Institute of Forensic Sciences, Gujarat Forensic Sciences University, Gandhinagar, India

A R T I C L E I N F O A B S T R A C T

Keywords: The non-metric characteristics like the tooth shape, number of cusps and other structural features contribute for the
Dental anthropology forensic and anthropological investigations and in establishing the population inter-relationships. The mandibular
Hardy-Weinberg equilibrium 2nd premolar and the maxillary 2nd molars often show variations in their occlusal morphology. The Hardy-
Population genetics
Weinberg equilibrium is applied in a novel way to determine the cusp phenotype in these two teeth in a
Cusp pattern
representative sample from Gujarati population. The dental stone models of 295 students of dental college in the age
Dental non metric trait
range of 19–23 years were used for this study. The two-cusped second premolar was present in around 60% the
subjects on both the right and left side. There is no significant difference in the distribution of the number of cusps in
both the right and left second premolars between male and female subjects. There was a significant difference in the
distribution of the number of cusps in left second molar between male and female (x2 = 4.073, df = 2, p < 0.05). The
strongest right - left correlation in the number of cusps was observed in maxillary 2nd molar (r = 0.809, p < 0.001).
On application of Hardy-Weinberg equilibrium, it was found that there was variation in the observed frequencies
and in the expected frequencies for the cusp number phenotypes in both the teeth.

Introduction Hardy – Weinberg equilibrium

Dental morphological traits have been extensively studied to assess
morphological affinities and variations among different population
groups [1,2]. These traits often appear discrete and heritable and show
different degrees of expression [3,4]. Such variations in the dental traits
are significantly correlated with the genetic and geographic factors [5].
In population genetics, the phenotypic characteristics in teeth are
closely related to the genotype and this ‘phenogenetic’ relationship
favours the prediction of a particular phenotype without a need for
genetic analysis [6]. Also the phenogenetic drift can define how much
genotype can evolve without a corresponding change in phenotype
[3,6]. The genetic information not only specifies the particular
characteristics of a tooth, but also determines the inheritance pattern
and the population variations of that trait. The environmental factors
also tend to play a role in the phenotypic manifestation of dental traits;
however this has not yet been fully quantified [7,8]. The degree of
variability of majority of dental non metric traits is between 0.4 to 0.9
[9]. The frequency of occurrence of each trait is phenotypic and the
genotype frequency is unknown however, some genes act pleiotropi-
cally on multiple dental phenotypes [10].
The Hardy –Weinberg (H-W) equilibrium is a principle which states
that alleles and genotype frequencies in a population will remain constant
from generation to generation in the absence of other evolutionary
influences. It is a mathematical equation derived by Geoffrey H. Hardy
(1877–1947), a mathematician and Wilhelm Weinberg (1862–1937), a
Physician [11]. In the field of population genetics, the H-W equilibrium is
considered as a model to estimate the frequencies of alleles and genotypes
[12]. Allele is a variant of a gene and for one gene there could be 2 alleles
(one from mother and one from father). Allele frequency is nothing but
the number of times that particular allele is present in a particular loci of
chromosome. It is determined by counting how many times the allele
appears in the population then dividing by the total number of copies of
the gene. The frequency of a particular allele in a population will always
remain constant, if there is no evolutionary change in that population.
Once the allele frequency is known, it is possible to estimate the genotype
frequency using the H-W principle. The allele frequency or the genotype
frequency will remain same for a particular gene from generation to
generation when the H-W equilibrium is met. In a simple genetic locus
there are two alleles A and a. The Hardy-Weinberg equation is expressed

* Corresponding author.
E-mail addresses: jppillaigdch@gmail.com (J.P. Pillai), dhwani.patel@gfsu.edu.in (D. Patel).

http://doi.org/10.1016/j.fsir.2019.100046
Received 18 July 2019; Received in revised form 3 October 2019; Accepted 6 October 2019
Available online 17 October 2019
2665-9107/© 2019 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-
J.P. Pillai, D. Patel
as p2 + 2pq + q2 = 1, where p is the frequency of the "A" allele and q is the
frequency of the "a" allele in the population. In the above mentioned H-W
equation, p2 represents the frequency of the homozygous genotype AA, q2
represents the frequency of the homozygous genotype aa, and 2pq
represents the frequency of the heterozygous genotype Aa. Also the sum of
the allele frequencies for all the alleles at the locus is 1, and hence
p + q = 1 [13]. The H-W equation is used to calculate the frequencies of
three genotypes when the p and q allele frequencies are known. The total
number of possible genotypes is calculated using the formula: G = n
(n+1)/2, where ‘n’ is the number of different alleles of a particular gene
and the Hardy-Weinberg equation can be used to measure whether the
observed genotype frequencies in a population differ from the frequencies
predicted by the equation.
Dental non metric traits
Dental morphology is highly genetically controlled and the non-
metric dental traits have been used in modern contexts to distinguish
between populations [14]. It was not until 1991 that the standards for
some non-metric dental traits were established [15]. The cusp size,
number, location and occlusal groove patterns are considered as some of
the main dental crown morphological traits in the field of dental
anthropology. Its mode of inheritance and the diversity in different
populations are being studied widely. In the permanent dentition, the
mandibular 2nd premolar and maxillary 2nd molar most often show
variations in number of cusps. In the former there are two cusp (2C) and
three cusp (3C) variants and in the latter there are three cusp (3C) and four
cusp (4C) variants [16]. The scoring of cusp number traits using the
reference plaque is easy and the developmental chronology of these two
teeth is almost similar in humans [17]. The dental non-metric
characteristics are more genetically related and because of the evolution
of the population genetics, a link between the genotype and the
phenotype, what is known as “phenogenetics” relationship could be
established [6]. The phenotypic variations in the tooth also give valuable
information about the migration pattern in humans and in understanding
the phylogenetic relationships between different populations in addition
to establishing the hereditary link. In a novel way, if we consider the cusp
number as the ‘allele’ and the right – left combination of the cusp type as
the ‘phenotype’, then the possible phenotypes may be calculated similar
to the above genotypes calculation. For example, if there are two types of
premolars based on the number of cusps (2C and 3C) then the possible
right and left combinations are : 2(2 + 1)/2 = 3. i.e., 2C-2C; 3C-3C; and
2/3C-3/2C combinations. The phenotype is said to be symmetrical if
both right and left shows same number of cusps (Homocusped) and
asymmetrical if the number differs (Heterocusped). The goal of this present
study was to propose a novel method applying the Hardy-Weinberg
equilibrium to study the distribution of cusp traits in teeth. To the best of
our knowledge, there are no previous studies in the literature that have
applied the Hardy-Weinberg equilibrium to study the dental cup
phenotype variations in the Indian population.
Study population
The study population was a representative sample of an ethnic group
traditionally from Gujarat, a state in the north western part of the Indian
peninsula, that speak Gujarati, an Indo-Aryan language. As the subjects
were selected from the premier Government dental institute in the state of
Gujarat, they represented the population of diverse communities in
Gujarat.
Materials and method
The upper and lower dental models of 295 dental students in the age
group of 19 to 23 years with a mean age of 21.03 years  1.31 were
selected. Institutional ethical committee’s approval was obtained prior to
the start of the study. The inclusion criteria included those casts with good
2
FSIR 1 (2019) 100046
anatomic details especially of the mandibular 2nd premolar and the
maxillary 2nd molars. The teeth with restorations or with crown and
bridges were excluded. The study subjects included 140 males and 155
female originating from different parts of Gujarat state and with diverse
religious and caste backgrounds. Informed signed consents were obtained
from the subjects prior to the start of the impression procedures. Using a
magnifier glass and the Arizona State University Dental Anthropology
System (ASUDAS) plaques as standard, the number of cusps in
mandibular 2nd premolar and in maxillary 2nd molar on both the sides
were counted and noted in the performa. The collection of the
quantitative data began with the operational definition of the cup trait
expression. The definition in the present study focused on the number of
cusp elevations present on the buccal and the lingual halves of the
occlusal surfaces of mandibular 2nd premolar using ASUDAS plaques as
reference. Similarly, the hypocone (distolingual - 4th cusp) expression in
the maxillary 2nd molar was observed and based on its presence or
absence, the cusp number was considered as 4 or 3 respectively. The
quantitative analysis of the cusps was done twice on all the samples on
two different occasions by the same observer. The data was stored in the
Excel sheet and the statistical analysis was performed using SPSS version
23.0 (SPSS Inc, Chicago, IL).
Calibration
The author practiced the handling of ASUDAS plaque by making
repeated series of observations with a gap of one month between the first
and the second recording. The recorded scorings were compared and this
repeated comparisons led to standardization of concepts.
Statistical analysis
The intra-observer agreement between the first and second recording
of data was tested using Cohen’s Kappa score. The extent of the
association between the number of cusps and gender was assessed using
chi-square analysis. The correlation study was performed using Spearman
two-tailed, to test the inter-trait and intra-trait variations. The
results were interpreted as follows: no correlation if |r| < 0.3, correlation
if 0.3 < |r| < 0.5, and strong correlation if 0.5 < |r| < 1 [18]. The paired
t-test was applied to check the difference in the frequency distribution of
number of cusps in the study teeth between right and left side. The
significance level was set to P < 0.05
Results
Intra-observer error
The Cohen’s Kappa score for the intra-observer agreement for the
quantification of cusps number trait in both the teeth was above 0.70.
(Table 1)
Frequency distribution of number of cusps in mandibular 2nd premolar
Of the total individuals studied, around 60% had 2 cusp mandibular
2nd premolars. The 2 cusp premolars were almost equally distributed in
both the right and left side for both genders. The females had higher
frequency distribution of 2 cusp type than males; however the difference
was not statistically significant. (Table 2) The males had higher frequency
Table 1
Intra-observer error Cohen Kappa results.
Non Metric-Trait Tooth (FDI system) Kappa score
(ASUDAS plaque)
P/2 CUSP Mandibular right 2nd Premolar(# 35) 0.762
Mandibular left 2nd Premolar (# 45) 0.714
HYPOCONE –CUSP 4 Maxillary right 2nd Molar(# 17) 0.792
Maxillary left 2nd Molar (# 27) 0.801
J.P. Pillai, D. Patel FSIR 1 (2019) 100046

Table 2
Table showing the frequency distribution of the number of cusps in mandibular second premolars and in maxillary second molars and their association with gender.
Tooth (FDI system) No. of cusps Sex x2
Male Female Total

Count % Count % Count % Value df Sig*.

Mandibular right 2nd Premolar(# 35) 2 84 60.0% 95 61.3% 179 60.7% 0.051 1 0.821
3 56 40.0% 60 38.7% 116 39.3%
Mandibular left 2nd Premolar (# 45) 2 81 57.9% 96 61.9% 177 60.0% 0.510 0.475
3 59 42.1% 59 38.1% 118 40.0%
Maxillary right 2nd Molar(# 17) 3 48 34.3% 60 38.7% 108 36.6% 0.620 0.431
4 92 65.7% 95 61.3% 187 63.4%
Maxillary left 2nd Molar (# 27) 3 42 30.0% 64 41.3% 106 35.9% 4.073 0.044
4 98 70.0% 91 58.7% 189 64.1%
*
Significant at P < 0.05.

distribution of 3 cusp type than females, however the difference was not
statistically significant.
Frequency distribution of number of cusps in maxillary 2nd molar
The 4 cusp type was the most predominant type in this tooth. More than
63% of the study models demonstrated 4 cusp type on both the sides.
(Table 2) The 4 cusp type 2nd molar was more predominant in male than
female on both sides. The gender difference in the distribution of the number
of cusps was significant only on the left side (#27) and not on the right (#17).
71.2% were having bilateral symmetry in cusp number in both the teeth.
Overall there was only 14.7% significant correlation in the distribution of
symmetrical and asymmetrical cusp numbers in both the teeth.
Paired t-test for right –left distribution of number of cusps
On the overall sample, there was no significant difference in the
distribution of cusp number in the antimeres. However, in male samples
there was significant difference in the distribution of cusp numbers
between #17 and #27. (Table 4)

Right-left correlation in the number of cusps Cross-tabulation of cusp phenotypes

Female had higher % of intra-trait correlation (r = 0.58, p < 0.05) in
the number of cusps between #35 and #45 than the males (r = 0.45,
p < 0.01). The highest intra-trait correlation in the number of cusps was
found between #17 and #27 in males (r = 0.873, p < 0.01). A significant
weak inter-trait correlation in the number of cusps was observed between
#35 and #17 in female (r = 0.19; p < 0.05). The difference in the number
of cusps between right and left side among mandibular 2nd premolar and
maxillary 2nd molar was significant in the latter (Table 3).
Symmetry/asymmetry in cusp number between sides
In the overall sample there was no significant difference in the number
of cusps between right and left side in the study teeth. Out of the total
sample, 76.9% were having symmetrical distribution of the number of
cusps in mandibular 2nd premolar and 91.2% in maxillary 2nd molar and
Table 5 shows the results of the inter-trait cusp phenotype
combinations between mandibular 2nd premolar and maxillary 2nd
molar. It reveals a weak positive correlation (r = 0.147; p < 0.001) in the
phenotype combinations between the two non- metric traits. In the
overall sample, 46.2% showed ‘homocusped’ pattern in both the teeth.
Novel application of Hardy-Weinberg equilibrium
The novelty in this study was that the right and left phenotype combi-
nation of teeth was modelled as allele combination, just like in geneotype.
The cusp frequencies correspond to the allele frequency and the right –left
combination of the cusp number is considered as the genotype as in actual
H-W equilibrium. Based on the observed frequencies of cusp phenotype, the
expected frequencies of the number of cusps and cusp phenotype combi-
nations were calculated by applying the H-W equation in a novel method.

Table 3
Table showing the non parametric correlation between number of cusps between the teeth of interest in male and female subjects.
Correlation between teeth Male Female Total
(FDI system)
Correlation Coefficient (R) Sig*. Correlation Coefficient (R) Sig*. Correlation Coefficient (R) Sig*.

#45 & #35 0.455 0.000 0.577 0.000 0.518 0.000

#17 & #27 0.873 0.000 0.759 0.000 0.809 0.000
#45 & #17 0.190 0.024 0.305 0.000 0.252 0.000
#45 & #27 0.185 0.029 0.236 0.000 0.212 0.000
#35 & # 27 0.275 0.001 0.253 0.002 0.265 0.000
#35 & #17 0.342 0.000 0.187 0.020 0.261 0.000
*
Correlation is significant at the 0.05 level (2-tailed).

Table 4
Table showing the results of the independent t-test between the right and left teeth.
Pair Male Female Total

Mean S.D. Sig*. Mean S.D. Sig*. Mean S.D. Sig*.

#45 - #35 0.021 0.515 0.624 0.006 0.448 0.858 0.007 0.48 0.809
#17 - #27 0.042 0.236 0.033 0.025 0.34 0.347 0.007 0.297 0.696
*
Difference is significant at 0.05 level (2-tailed).

3
J.P. Pillai, D. Patel FSIR 1 (2019) 100046

Table 5
Table showing the descriptive statistics of the cross tabulation of the cusp phenotype (right –left combination) in mandibular 2nd premolar and maxillary 2nd molar.
Mandibular 2nd Premolar Maxillary 2nd Molar (Rt.- Lt.) Total Correlation Co-efficient x2 Sig*.
(Rt. - Lt.)
3C-3C 4C-4C 3C-4C 4C-3C

Count % Count % Count % Count % Count %

2C-2C 63 21.4 68 23.1 8 2.7 5 1.7 144 48.8 0.147 34.08 0.000
3C-3C 11 3.7 68 23.1 2 0.7 2 0.7 83 28.1
2C-3C 10 3.4 22 7.5 2 0.7 1 0.3 35 11.9
3C-2C 10 3.4 17 5.8 2 0.7 4 1.4 33 11.2
Total 94 31.9 175 59.3 14 4.7 12 4.1 295 100
*
Significant at P < 0.05.

Estimating the cusp frequencies in Premolar. There is a marked variation on in the frequency distribution of the cusp
phenotype traits in the observed data and the expected data. A reverse
trend is expected in the ‘heterocusped’ phenotype in both the teeth and the
variation was maximum in maxillary 2nd molar ( 0.37)(Chart 1 ).

Discussion

In Dental anthropology, the dental traits helps the anthropologists to

understand inter and intra-population dynamics and variations with due
considerations to the geographical, environmental and socio-economic
parameters [19]. The present study examined the expression and
distribution pattern of two dental morphological traits in the Gujarati
population, which represents the ancestral North Indian gene and appear
much more diverse than other South Asian populations [20]. The non-
metric traits that were included in the study were the cusp numbers in
mandibular second premolar and maxillary second molars. The frequency
distribution of cusp traits in mandibular second premolar was 60% and
40% among two cusp and three cusp types respectively and the maxillary
second molar was predominantly of four cusp type. There is hardly any
single study comparing the cusp patterns in these two teeth. Two studies
from Kerala, a state in Southern India have analysed the frequency
distribution mandibular second premolar according to cusp numbers
[21,22]. One study has shown the distribution of two cusp and three cusp
mandibular second premolar distributions in 52.8% and 44.4%
respectively [21]. On the contrary the other study by BabyTK et al has
reported mandibular second premolar with more than one lingual cusp in
52.92% of the subjects. [22]. Their study observed one buccal cusp and
two lingual cusps (3 cusp type) and three lingual cusps (4 cusp type). The
present study did not observe the 4 cusp type premolars in the Gujarati
samples. The Kerala population represents the Dravidian ancestry and
phylogenetic analysis has shown clear cut genetic demarcation from
Gujarati population [23]. However, the distribution of the cusp pattern in
mandibular 2nd premolars is almost similar in both the Kerala and the
Gujarati population. There was a predominance of two cusp type
premolar (54.6%) in a study on Korean population, which was similar to
the results of the present study (60%). The three cusp mandibular 2nd
premolar was higher in Korean (45.5%) than in Caucasian (26.3%)
population. [24] The Caucasoid population include groups such as West
Asians (Arabs), Europeans, Northern Africans, and south Asians who are
closely related and can be distinguished from Mongoloid or Negroid
population [25]. The two cusp form (66.3%) was far more common than
the three cusp form (25.4%) in Singaporean Chinese population which
was consistent with the present study [26]. However the distribution of
three cusp form was higher in the present Gujarati population than the
Chinese population. Also the asymmetry in the cusp numbers was higher
(23.1%) in the present study when compared to the Singaporean Chinese
population (8.3%). The Singaporean Chinese are the descendants of the
Mongoloid race in mainland China and they constitute the largest ethnic
group (76.2%) of the population of Singapore. The shovelling of incisors
which is a characteristic feature in Mongoloid race has been extensively
studied [27]. According to Ludwig (1957), there was greater variation in
the distribution of the two cusp and three cusp premolars in the
Mongoloid race [28]. In his study the cusp distribution in the Caucasoid

4
J.P. Pillai, D. Patel
Chart 1. Chart showing the difference in the observed and expected frequencies of the cusp number phenotypes in both the teeth.
race was very much similar to the findings of the present study. The
predominant distribution of two cusp patterns over the three cusp pattern
premolar in the present study is consistent with the results from Kerala
(South Indian) [21], Korean [24], Singaporean Chinese [26] and
Peshawar [29] populations. However, the studies from Iranian [30],
Lahore [31], Dhaka [32], Jordanian [33] and Thai [34] populations have
revealed a reverse trend in the cusp trait distribution in mandibular
second premolar. The distribution of the three cusp pattern was dominant
in theses populations. The Thai tribal population from Northern Thailand
showed the maximum distribution of 3 cusp premolars (86.6%). Though
this population has the features of Mongoloid race, the premolar cusp
pattern predominantly featured three cusp form unlike the other
mongoloid population like in Singapore Chinese and Koreans [34].
The gene flow because of migration and mixture with native populations
or the local evolution without genetic influence could be attributed as
reasons for the variations in the tooth morphology [33]. Thus it is
noteworthy to mention that the expression of this trait reveals
considerable variations in the human population. The second trait
analysed in this study, the fourth cusp (Hypocone) in the maxillary second
molar is also thought to vary between populations. The distribution of this
trait is around 64% and in the remaining 36% the second molar was of
three cusp type. The three cusp type was found in 15.8% of the Davangere
population in Karnataka, South India [35]. Genetic study has shown
difference in the Karnataka’s Dravidian population and the Gujarat’s
Indo-Aryan population [23]. Studies have shown relationship between
the expression of the maxillary second molar hypocone and the first
molar hypocone and also with the fifth cusp (cusp of Carabeli) in
maxillary first molar [35–38]. The present study compared the expression
of the hypocone in the upper second molar with the lingual cusp trait in
the lower second premolars and found a significant weak positive inter-
trait correlation. The hypocone reduction or three cusp type maxillary
second molar was observed in around 29% of the Jordanian population
[36] which is around 36% in the present study. In consistent with the
findings of the Jordan population and the present Gujarati population, the
Thai and the Aka tribal groups in Northern Thailand also showed
predominant expression of the fourth cusp in maxillary second molar
[33]. Unlike the other studies on cusp traits, the present study applied the
frequency distribution of the cusp pattern in the present sample using
H-W equilibrium to calculate the expected frequencies of the traits in the
same population. An earlier study by Cucina proposed a new approach to
the estimation of intra-site variability of dental morphological traits when
they are dichotomized into presence vs absence [13]. The present study,
like the study by Cucina had adapted the genetic rules to phenotypic
features by considering the morphological (cusp) frequencies to allele
frequency. But the present study extended in a novel way by considering
the right and left combinations of cusp frequency in the antimeres for
genotype.
5
FSIR 1 (2019) 100046
Conclusion
A novel approach of applying the Hardy-Weinberg principle in a cusp
phenotype analysis is reported here in a representative population of
Gujarat with diverse religious and community profiles. Such dental
anthropological studies can provide valid information about the
variations of a particular trait in a given population. The authors propose
the smaller sample size, the possible genetic drift and the gene flow as
possible reasons for the variations in the observed phenotype frequency
and the expected phenotype frequencies in the present sample with
religious and caste diversity. However, large sample studies and
inheritance studies on these traits need to be designed for further
validation.
Financial disclosure
This research did not receive any specific grant from funding agencies
in the public, commercial, or not-for-profit sectors.
Declaration of Competing Interest
None.
Acknowledgements
The authors wish to acknowledge Dr. Bhargav Patel, Assistant
Professor, IFS, GFSU for his guidance and technical support for this
research and Dr. Asaithambi Balamurugan, Forensic Odontologist,
Chennai for providing the ASUDAS plaques. The author also acknowl-
edges all the Dental students of GDCH Ahmedabad who volunteered to
provide their dental models for this study.
References
[1] G. Richard Scott, Christy G. Turner, Anthropology of Modern Human Teeth,
Cambridge University Press, Cambridge, 1997.
[2] R. Vargiu, A. Cucina, A. Coppa, Italian populations during the Copper Age: assessment
of biological affinities through morphological dental traits, (Hum. Biol. 81 (2009) 91–
104.
[3] J. Jernvall, H.S. Jung, Genotype, phenotype, and developmental biology of molar
tooth characters, (Yearb. Phys. Anthropol. 43 (2000) 171–190.
[4] C.S. Larsen, M.A. Kelley, Introduction, in: M.A. Kelley, C.S. Larsen (Eds.), Advances in
Dental Anthropology, Wiley-Liss, New York, 1991, pp. 1–5.
[5] C.A. Brewer-Carias, S. LeBlanc, J.V. Neel, Genetic structure of a tribal population, the
Yanomama Indians. XII. Dental microdifferentiation, (Am. J. Phys. Anthropol. 44
(1976) 5–14.
[6] K.M. Weiss, S.M. Fullerton, Phenogenetic drift and the evolution of genotype-
phenotype relationships, (Theor. Popul. Biol. 57 (2000) 187–195.
[7] G.C. Townsend, P. Dempsey, T. Braun, G. Kaidonis, L. Richards, Teeth, genes and the
environment, (Persp. Hum. Biol. 4 (1994) 35–46.
J.P. Pillai, D. Patel
[8] G.C. Townsend, T. Hughes, M. Luciano, M. Bockmann, A. Brook, Genetic and
environmental influences on human dental variation: a critical evaluation of studies
involving twins, (Arch. Oral Biol. 54 (Suppl) (2009) S45–S51.
[9] T. Hanihara, Morphological variation of major human populations based on
nonmetric dental traits, (Am. J. Phys. Anthropol. 136 (2008) 169–182.
[10] T. Hughes, G. Townsend, Gene for teeth – drawing inferences from family data, in: G.
Townsend, E. Kanazawa, H. Takayama (Eds.), New Directions in Dental Anthropology.
Paradigms Methodologies and Outcomes, The University of Adelaide Press, Adelaide,
2012, pp. 22–34.
[11] James F. Crow, "Hardy, Weinberg and language impediments", (Genetics 152 (3)
(1999) 821–825.
[12] P.H. Hedrick, Genetics of Population, Science Books International, Boston, 2000.
[13] A. Cucina, Intra-population dental morphological variability among the Prehispanic
Maya, (J. Comp. Human Biol. 67 (5) (2016) 384–396.
[14] Heather J.H. Edgar, Estimation of Ancestry using dental morphological
characteristics, (J. Forensic Sci. 58 (2013) S3–S8.
[15] Christy G. Turner II, Christian R. Nichol, G. Richrd Scott, Scoring procedures for key
morphological traits of the permanent dentition: the Arizona State university dental
anthropology system, in: Marc A. Kelley, Clark Spencer Larsen (Eds.), Advances in
Dental Anthropology, wiley-Liss, Inc., New York, 1991, pp. 13–31.
[16] C. Scheid Rickne, Woelfel’s Dental Anatomy, 8th ed., Lippincott Williams & Wilkins,
Philadelphia PA,19103 USA, 2012.
[17] M.M. Ash, S.J. Nelson, Wheeler’s Dental Anatomy, Physiology and Occlusion,
Saunders, Philadelphi, PA, 2003.
[18] J. Cohen, Statistical power analysis for the behavioral-sciences, (J. Percept. Motor
Skill. 67 (1988) 1007.
[19] R.S. Corruccini, R.H. Yap Potter, Developmental correlates of crown component
asymmetry and occlusal discrepancy, (Am. J. Phys. Anthropol. 55 (1981) 21–31.
[20] Marina Silva, Marisa Oliveira, Daniel Vieira, Andreia Brandão, Teresa Rito, Joana B.
Pereira, Ross M. Fraser, Bob Hudson, Francesca Gandini, Ceiridwen Edwards, Maria
Pala, John Koch, James F. Wilson, Luísa Pereira, Martin B. Richards, Pedro. Soares,
Genetic chronology for the Indian Subcontinent points to heavily sex-biased
dispersals, (BMC Evol. Biol. 17 (1) (2017) 88.
[21] S. Sunil, D. Gopakumar, Prevalence of the two variants of mandibular second
premolars in Kerala population, (Int. J. Odontostomat 6 (2012) 375–377.
[22] T.K. Baby, S. Sunil, S.S. Babu, Nonmetric traits of permanent posterior teeth in
Kerala population: a forensic overview, (J. Oral Maxillofac. Pathol. 21 (2017) 301–
308.
[23] Lekha Pandit, Anitha D’Cunha, Chaithra Malli, Genetic variations in the Dravidian
population of South West coast of India: implications in designing case-control studies,
(Indian J. Med. Res. 145 (6) (2017) 753–757.
FSIR 1 (2019) 100046
[24] Park H.-Y. Yoo-HI, S.-H. Kim, Occlusal surface analysis of mandibular premolars in
koreans, (Korean J. Phys. Anthropol. 28 (2015) 145–153.
[25] A. Pakstis, C. Gurkan, M. Dogan, H.E. Balkaya, S. Dogan, P. Neophytou, L. Cherni, S.
Boussetta, H. Khodjet-el-khil, N. Salvo, K. Janssen, G.H. Olsen, S. Hadi, E.
Almohammed, V. Pereira, D. Truelsen, O. Bulbul, U. Soundararajan, K. Kidd, Genetic
relationships of European, Mediterranean, and SW Asian populations using a panel of
55 AISNPs, (Eur. J. Hum. Genet. (2019) 1–6.
[26] H.S. Loh, Coronal morphology of the mandibular second premolar in the Singaporean
Chinese, (Aust. Dent. J. 38 (1993) 283–286.
[27] H. Yaacob, P. Nambiar, M.D.K. Naidu, Racial characteristics of human teeth with
special emphesis on the mongoloid dentition, (Malaysian J. Pathol. 18 (1) (1996) 1–7.
[28] F.J. Ludwig, The mandibular second premolars : morphologic variation and
inheritance, (J. Dent. Res. 36 (2) (1957) 263–273.
[29] D.B. Khan, Z.-Ur Rehman, F. Khan, H. Khan, Frequency of bicuspid and tricuspid form
of mandibular second premolar at Khyber College of Dentistry Peshawar, (J. Med. Sci.
21 (2013) 118–120.
[30] R. Mosharraf, F. Hajian, Occlusal morphology of the mandibular first and second
premolars in Iranian adolescents, (Dent. Anthropol. J. 17 (3) (2004) 94–96.
[31] A. Ahmed, N. Iqbal, M.Z. Baig, T. Ayub, Frequency of type traits of mandibular second
premolars — a study, (Pak. Oral Dent. J. 36 (2016) 430–432.
[32] M.R. Hossain, M.K. Alam, Z. Shawkat, S.I. Quasem, M.Z. Hossain, M.A. Islam, Cuspal
variation in mandibular second premolars, (Bangladesh Dent. Coll. J. (2009) 7–9.
[33] A. Khraisat, F. Alsoleihat, K. Subramani, M.A. Al-Rabab’ah, M.K. Al-Omiri, I. Abu-
Tahun, Multiple lingual cusps trait on mandibular premolars and hypoconulid
reduction trait on mandibular first molar in living jordanian population. intra- and
inter-trait interactions, (Coll. Antropol. 37 (3) (2013) 885–894.
[34] Y. Manabe, R. Ito, Y. Kitagawa, J. Oyamada, A. Rokutanda, S.Y. Nagamoto, S.
Kobayashi, K. Kato, Non-metric tooth crown traits of the that, Aka and Yao tribes of
Northern Thailand, (Arch. Oral Biol. 42 (4) (1997) 282–291.
[35] P. Poornima, M. Krithiga, S. Sasalwad, N.B. Nagaveni, Prevalence of few variant dental
features in children aged 11-16 years in Davangere, a city in Karnataka, (J. Forensic
Dent. Sci. 8 (2016) 13–17.
[36] A. Khraisat, F. Alsoleihat, K. Subramani, S.T. Taha, Z.B. Al-Rabab’ah MA Al-Bitar,
Hypocone reduction and Carabelli’s traits in contemporary jordanians and the
association between Carabelli’s trait and the dimensions of the maxillary first
permanent molar, (Coll. Antropol. 35 (1) (2011) 73–78.
[37] H.J. Keene, The relationship between Carabeli’s trait and the size, number and
morphology of maxillary molars, (Arch. Biol. 13 (1968) 1023–1025.
[38] M. Takahashi, S. Kondo, G.C. Townsend, E. Kanazawa, Variability in cusp size in
human molars with particular reference to Hypocone, (Arch. Oral Biol. 52 (2007)
1146–1154.